The origin(s) of extant amphibians: a review with emphasis on the "lepospondyl hypothesis"

The origins of the extant amphibians (frogs, salamanders, caecilians) remain controversial after over a century of debate. Three groups of hypotheses persist in the current literature: the “temnospondyl hypothesis” (TH) which roots Lissamphibia Haeckel, 1866 (the smallest clade composed of the extant amphibians) within the Paleozoic temnospondyls, the “lepospondyl hypothesis” (LH) which postulates a monophyletic Lissamphibia nested within the Paleozoic lepospondyls, and the “polyphyly hypothesis” (PH), according to which the frogs and the salamanders are temnospondyls while the caecilians are lepospondyls. The discovery of the Middle Jurassic to Pliocene albanerpetontids, which are very similar to the extant amphibians, has complicated rather than resolved this situation. We present a review of recent publications and theses in this field, several of which show more support for the LH than for the TH and considerably more than for the PH. In addition, we show that there is no particular attraction between long-bodied lissamphibians (caecilians) and long-bodied lepospondyls (such as the lysorophians): when they are removed from two published matrices, reanalyses nonetheless find the LH. In one case the LH is found even when all salamanders are removed as well. We furthermore propose that the complex of characters called the salamander mode of autopodium development is (in its less extreme forms) plesiomorphic for limbed vertebrates, so the apparent presence of this mode of development in temnospondyls cannot support the TH or the PH. Still, a consensus will not be reached soon, despite the increasing range of data and types of analysis that are used (morphological, molecular and combined phylogenetics, development biology, molecular divergence dating, paleontological supertree dating, combined dating, and calculation of confidence intervals on first appearances in the fossil record). We present examples of pertinent character state distributions and explore a large gap in the fossil record of small stegocephalians.